大刍草稀有等位基因促进玉米密植高产

密植是提高作物单位面积产量、促进粮食增产的重要途径之一。叶夹角是影响玉米(Zea mays)密植的关键因子。中国农业大学田丰课题组最近克隆了2个调控玉米叶夹角的数量性状位点(QTL)——UPA1和UPA2, 揭示了这2个位点的功能基因(brd1和ZmRAVL1)通过油菜素内酯(BR)信号通路调控叶夹角。UPA2位于ZmRAVL1上游9.5 kb, 可与DRL1蛋白结合。另一个影响玉米叶夹角的蛋白LG1可以激活ZmRAVL1的表达; DRL1蛋白与LG1蛋白直接互作抑制LG1对ZmRAVL1的激活表达。玉米祖先种大刍草(teosinte)的UPA2位点序列与DRL1蛋白结合能力更强, 导致大刍草ZmRAVL1的表达受到更强的抑制, 下调表达的ZmRAVL1进一步使下游基因brd1的表达下调, 进而降低叶环区的内源BR水平, 导致叶夹角变小。将大刍草的UPA2等位基因导入到玉米中或对玉米中ZmRAVL1进行基因编辑, 在密植条件下均可显著提高玉米产量。上述发现为高产玉米品种的分子育种改良提供了重要理论基础和基因资源。     

大刍草稀有等位基因促进玉米密植高产

密植是提高作物单位面积产量、促进粮食增产的重要途径之一。叶夹角是影响玉米(Zea mays)密植的关键因子。中国农业大学田丰课题组最近克隆了2个调控玉米叶夹角的数量性状位点(QTL)——UPA1和UPA2, 揭示了这2个位点的功能基因(brd1和ZmRAVL1)通过油菜素内酯(BR)信号通路调控叶夹角。UPA2位于ZmRAVL1上游9.5 kb, 可与DRL1蛋白结合。另一个影响玉米叶夹角的蛋白LG1可以激活ZmRAVL1的表达; DRL1蛋白与LG1蛋白直接互作抑制LG1对ZmRAVL1的激活表达。玉米祖先种大刍草(teosinte)的UPA2位点序列与DRL1蛋白结合能力更强, 导致大刍草ZmRAVL1的表达受到更强的抑制, 下调表达的ZmRAVL1进一步使下游基因brd1的表达下调, 进而降低叶环区的内源BR水平, 导致叶夹角变小。将大刍草的UPA2等位基因导入到玉米中或对玉米中ZmRAVL1进行基因编辑, 在密植条件下均可显著提高玉米产量。上述发现为高产玉米品种的分子育种改良提供了重要理论基础和基因资源。     

玉米叶夹角形成的遗传基础与分子机制解析*

玉米产量取决于植株捕获光能和固定CO₂合成有机化合物的效率。叶夹角是株型重要性状之一,较小叶夹角有利于提高玉米植株光合作用效率和种植密度,因而有利于提高玉米产量。研究表明玉米叶夹角为多基因控制的复杂数量性状,其遗传力较高,主要受基因的加性效应调控。目前,利用数量性状位点(quantitative trait loci, QTL)定位和全基因组关联分析(genome-wide association study, GWAS)等方法已鉴定数百个玉米叶夹角相关QTL;结合突变体分析等方法,已克隆数十个调控叶夹角关键基因,这为了解玉米叶夹角遗传机制提供了重要参考。由于前人研究所采用群体、分析方法及参考基因组版本不同,各研究之间所鉴定QTL差异较大,因此无法客观揭示叶夹角性状的遗传规律。为此,通过总结前人所定位叶夹角相关QTL和单核苷酸多态性(single nucleotide polymorphism,SNP)位点并构建一致性图谱,鉴定出叶夹角性状定位热点区间,并对调控叶夹角的已知基因进行功能分类。这不仅为了解玉米叶夹角的遗传结构、推动叶夹角相关重要基因克隆提供数据支撑,也对进一步开发叶夹角相关分子标记,指导玉米分子育种和提高玉米产量提供有益指导。     

两个荔枝品种的叶面积与叶重量估测模型

以‘黑叶’和‘怀枝’2个荔枝品种的成熟叶片为材料,对叶周长、叶长、叶宽、叶长宽乘积与叶面积、叶重量之间的相关性进行回归分析。结果表明,‘黑叶’和‘怀枝’均可通过叶周长、叶长、叶宽和叶长宽乘积进行叶面积估算,其中以叶长宽乘积的估算效果最好;‘黑叶’可通过叶周长、叶长、叶宽、叶长宽乘积和叶面积估算叶重量,其中以叶长指标的估算效果最佳;‘怀枝’基于上述5个叶形指标进行的叶重量估算效果不佳。     

羽衣甘蓝裂叶相关性状遗传分析

以羽衣甘蓝圆叶自交系‘0835’和裂叶自交系‘0819’为亲本,调查P1、P2、F1、F2群体莲座期4个裂叶相关性状表型数据,运用‘四世代主基因+多基因’遗传模型,对叶长、叶宽、叶形指数、叶缘缺刻数4个叶形相关性状进行遗传分析,探讨羽衣甘蓝裂叶相关性状的遗传规律,为羽衣甘蓝裂叶性状遗传、QTL定位及新品种选育奠定基础。结果表明:(1)4个性状均存在一定的杂种优势,其中叶缘缺刻数中亲优势达显著水平,4个性状均存在负向超亲优势。(2)叶长和叶宽均符合E-4模型,即由2对等加性主基因+加性-显性多基因共同控制;叶长主基因遗传率为83.80%,多基因遗传率为1.05%;叶宽主基因遗传率为22.28%,多基因遗传率为61.92%。(3)叶形指数和叶缘缺刻数均符合E-1模型,即由2对加性-显性-上位性主基因+加性-显性多基因控制;叶形指数主基因遗传率为93.73%,多基因遗传率为2.59%;叶缘缺刻数主基因决定了表型变异的91.18%。     

早实核桃叶片性状遗传规律研究

为了探索核桃叶片性状的遗传规律,为杂交后代性状早期选择提供依据,以2个正反交组合374个杂交单株7年生实生树为试材,测定了各正反交单株的叶长、叶宽、叶面积、叶形指数和比叶重等。结果表明:各性状的变异系数均在6%以上,比叶重的变异系数最高,在15%以上;叶形指数的遗传传递力在4个杂交组合中均大于100%;叶宽和叶形指数较其他3个性状的广义遗传力高,均在79%以上;叶面积的次数分布呈连续性变异,不同杂交组合正反交后代叶面积大小的分布主要集中在55~60 cm2,小叶型、中间型和大叶型植株在杂交后代群体中所占比率分别为13%~15%、74%~80%和10%以下;在绿岭、辽宁1号的正反交组合中,叶长、叶宽、叶面积、叶形指数和比叶重分别为12.52 cm、6.72 cm、55.27 cm2、1.88、53.18 g/m2和12.86 cm、6.71 cm、56.43 cm2、1.91、59.09 g/m2,反交后代的叶面积和比叶重2个性状均极显著大于正交后代;在绿岭、绿早的正反交组合中,各性状的值分别为12.63 cm、6.73 cm、57.25 cm2、1.94、63.54 g/m2和12.58 cm、6.80 cm、57.59 cm2、1.88、62.28 g/m2,正反交的叶长、叶宽、叶面积和比叶重性状均没有显著差异,正交后代的叶形指数显著大于反交后代。     

银杏叶片形态研究

通过对银杏不同枝类、不同叶序的叶片形态进行调查,结果表明:银杏不同枝类的叶片形态差异较大,叶宽、叶柄长、叶基角:多年生鳞枝＞一年生鳞枝＞一年生长枝;叶长、叶形指数:一年生长枝＞一年生鳞枝＞多年生鳞枝;叶面积:一年生和多年生鳞枝＞一年生长枝;有缺刻叶比例:一年生长枝＞多年生鳞枝＞一年生鳞枝.一年生长枝叶片的叶宽、叶长、叶面积、叶柄长、叶基角均随叶序的增加逐渐减小,叶形指数和有缺刻叶的比例则增加.一年生和多年生鳞枝1～5叶的叶长、叶宽和叶面积随叶序增加而逐渐增加,第5～6叶达最大,以后随叶序增加而逐渐减小,叶形指数和叶柄长度随叶序增加而增加,叶基角随叶序增加而减小.一年生长枝的第2叶、一年和多年生鳞枝的第4叶可作为品种描述的标准叶.     

大白菜部分形态性状的QTL定位与分析

应用352个标记位点的大白菜AFLP和RAPD图谱和一套栽培品种间杂交获得的重组自交系群体,采用复合区间作图的方法对大白菜9个形态性状进行QTL定位及遗传效应研究。在14个连锁群上检测到50个QTL:其中控制株型的QTL有5个;控制株高的QTL有6个;控制开展度的QTL有5个;控制最大叶长的QTL有7个;控制最大叶宽的QTL有4个;控制叶形指数的QTL有6个;控制中肋长的QTL有7个;控制中肋宽的QTL有4个;控制抽苔的QTL有6个。另外,估算了单个QTL的遗传贡献率和加性效应。这将为大白菜品种改良中形态性状的分子标记辅助选择提供理论依据。     

玉米杂交种掖单13号的SSR连锁图谱构建与叶夹角和叶向值的QTL定位与分析

为了增加单位面积产量, 玉米育种者已经开始了更密植更紧凑株型的选育。叶夹角和叶向值是评价玉米株型的重要指标。本研究以掖478×丹340的500个F2单株为作图群体, 构建了具有138个位点的SSR标记连锁图谱, 图谱总长度为1 394.9 cM, 平均间距10.1 cM。利用397个F2：3家系对叶夹角和叶向值进行QTL定位分析, 结果表明: 叶夹角和叶向值分别检测到6和8个QTL, 累计解释表型变异41.0%和60.8%, 单个QTL的贡献率在2.9%~13.6%之间。与叶夹角和叶向值有关的基因主要作用方式为加性和部分显性。此外两个性状共检测到9对上位性互作位点, 表明上位性互作在叶夹角和叶向值的遗传中也起较重要的作用。     

基于元分析的抗玉米丝黑穗病QTL比较定位

以玉米遗传连锁图谱IBM2 2005 Neighbors为参考图谱,通过映射整合不同试验中的抗玉米丝黑穗病QTL,构建QTL综合图谱。在国内外种质中,共发现22个抗病QTL,分布在除第7染色体外的9条玉米染色体上。采用元分析技术,获得2个“一致性”抗病QTL,图距分别为8.79 cM和18.92cM。从MaizeGDB网站下载“一致性”QTL区间内基因和标记的原始序列;采用NCBI网站在线软件BLASTx通过同源比对在2个“一致性”QTL区间内初步获得4个抗病位置候选基因。借助比较基因电子定位策略,将69个水稻和玉米抗性基因定位于玉米IBM2图谱上,在2个“一致性”QTL区间内分别发现1个水稻抗性基因,初步推断为抗病位置候选基因。本文结果为抗玉米丝黑穗病QTL精细定位和分子育种提供了基础。     

Integrated multiple population analysis of leaf architecture traits in maize (Zea mays L.)

Leaf morphology in maize is regulated by developmental patterning along three axes: proximodistal, mediolateral, and adaxial-abaxial. Maize contains homologues of many genes identified as regulators of leaf development in other species, but their relationship to the natural variation of leaf shape remains unknown. In this study, quantitative trait loci (QTLs) for leaf angle, leaf orientation value, leaf length, and leaf width were mapped by a total of 256 F(2:3) families evaluated in three environments. Meta-analysis was used to integrate genetic maps and detect QTLs across several independent QTL studies, on the basis of the previously reported experimental results for leaf architecture traits. Candidate gene sequences for leaf architecture were mapped in the integrated consensus genetic map. In total, 21 QTLs and 17 meta-QTLs (mQTLs) were detected. Among these QTLs, qLA1-1 and qLA2 were consistently detected in five and three populations respectively, and six of seven QTLs with contributions (R(2)) >10% were integrated in mQTLs. Six key mQTLs (mQTL1-1, mQTL2-1, mQTL3-3, mQTL5-1, mQTL7-2, and mQTL8-1) with R(2) of some initial QTLs >10% included 4-6 initial QTLs associated with 2-4 traits. Therefore, the chromosome regions for six mQTLs with high QTL co-localization might be hot spots of the important QTLs for the associated traits. Fifteen key candidate genes controlling leaf architecture traits coincided with 11 corresponding mQTLs, namely DWARF4, KAN3, liguleless1, TAC1, ROT3, AS2/liguleless2, PFL2, yabby9/SE/LIC/yabby15, mwp1, CYCD3;2, and CYCB1. In particular, DWARF4, liguleless1, AS2/liguleless2, yabby9/SE/LIC/yabby15, and CYCD3;2 were mapped within the important mQTL1-1, mQTL2-1, mQTL3-3, mQTL5-1, and mQTL7-2 intervals, respectively. Fine mapping or construction of single chromosome segment lines for genetic regions of these five mQTLs is worth further study and could be put to use in marker-assisted breeding. In conclusion, the results provide useful information for further research and help to reveal the molecular mechanisms with regard to leaf architecture traits.     

水稻叶片水势的QTL定位与候选基因分析

为探究叶片水势(LWP)相关基因在水稻(Oryza sativa)抗旱中的作用及其遗传机制,以热研2号(Nekken2)和华占(HZ)为亲本以及构建的120个重组自交系(RILs)群体为实验材料,对水稻分蘖期叶片水势进行检测,并利用前期基于高通量测序构建的分子遗传连锁图谱进行数量性状基因座(QTL)分析。结果表明,共检...     

Genetic Analysis and Major Quantitative Trait Locus Mapping of Leaf Widths at Different Positions in Multiple Populations

Background

Leaf width is an important agricultural trait in maize. Leaf development is dependent on cell proliferation and expansion, and these processes exhibit polarity with respect to the longitudinal and transverse axes of the leaf. However, the molecular mechanism of the genetic control of seed vigor remains unknown in maize, and a better understanding of this mechanism is required.

Methodology/Principal Findings

To reveal the genetic architecture of leaf width, a comprehensive evaluation using four RIL populations was performed, followed by a meta-analysis. Forty-six QTLs associated with the widths of leaves at different positions above the uppermost ear were detected in the four RIL populations in three environments. The individual effects of the QTLs ranged from 4.33% to 18.01% of the observed phenotypic variation, with 14 QTLs showing effects of over 10%. We identified three common QTLs associated with leaf width at all of the examined positions, in addition to one common QTL associated with leaf width at three of the positions and six common QTLs associated with leaf width at two of the positions. The results indicate that leaf width at different leaf positions may be affected by one QTL or several of the same QTLs. Such traits may also be regulated by many different QTLs. Thirty-one of the forty-six initial QTLs were integrated into eight mQTLs through a meta-analysis, and 10 of the 14 initial QTLs presenting an R2>10% were integrated into six mQTLs.

Conclusions/Significance

mQTL1-2, mQTL3-1, mQTL7, and mQTL8 were composed of the initial QTLs showing an R2>10% and included four to six of the initial QTLs that were associated with two to four positions in a single population. Therefore, these four chromosome regions may be hot spots for important QTLs for these traits. Thus, they warrant further studies and may be useful for marker-assisted breeding.     

Detection and integration of quantitative trait loci for grain yield components and oil content in two connected recombinant inbred line populations of high-oil maize

Improvement in grain yield is an important objective in high-oil maize breeding. In this study, one high-oil maize inbred was crossed with two normal maize inbreds to produce two connected recombinant inbred line (RIL) populations with 282 and 263 F_7:8 families, respectively. The field experiments were conducted under four environments, and eight grain yield components and grain oil content were evaluated. Two genetic linkage maps were constructed using 216 and 208 polymorphic SSR markers. Quantitative trait loci (QTL) were detected for all traits under each environment and in combined analysis. Meta-analysis was used to integrate genetic maps and detected QTL in both populations. A total of 199 QTL were detected, 122 in population 1 and 87 in population 2. Seven, 11 and 19 QTL showed consistency across five environments, across two RIL populations and with respective F_2:3 generations, respectively. 183 QTL were integrated in 28 meta-QTL (mQTL). QTL with contributions over 15% were consistently detected in 3–4 cases and integrated in mQTL. Each mQTL included 3–19 QTL related to 1–4 traits, reflecting remarkable QTL co-location for grain yield components and oil content. Further research and marker-assisted selection (MAS) should be concentrated on 37 consistent QTL and four genetic regions of mQTL with more than 10 QTL at bins 3.04–3.05, 7.02, 8.04–8.05 and 9.04–9.05. Near-isogenic lines for 100-grain-weight QTL at bin 7.02–7.03, for ear-length QTL at bin 7.02–7.03 and for rows-per-ear QTL at bin 3.08 are now in construction using MAS. Co-located candidate genes could facilitate the identification of candidate genes for grain yield in maize.     

ZmFBL41 Chang7-2: 玉米抗纹枯病的关键利器

由真菌Rhizoctonia solani引起的纹枯病严重危害玉米(Zea mays)和水稻(Oryza sativa)等作物的安全生产。R. solani的宿主范围广且抗源少, 加之相关的抗性机制研究有限, 导致纹枯病的危害长期得不到有效控制。近期, 中国科学家通过对318份玉米自交系进行全基因组关联分析, 筛选到1个与纹枯病抗性相关的、编码F-box结构域蛋白的候选基因ZmFBL41 (GRMZM2G109140)。ZmFBL41蛋白是SCF (SKP1-Cullin-F-box) E3泛素连接酶复合体的一员, 能介导复合体对肉桂醇脱氢酶ZmCAD的降解, 从而降低木质素的积累, 使玉米易感纹枯病。玉米抗病自交系Chang7-2中, 蛋白ZmFBL41 ^Chang7-2因2个关键氨基酸的变异, 不能结合并降解底物ZmCAD, 使木质素含量增加, 从而提高玉米对纹枯病的抗性。该研究率先揭示了SCF复合体可通过降解肉桂醇脱氢酶来调控植物免疫反应的新型分子机制, 为提高玉米及其它作物对纹枯病的抗性提供了重要理论依据和基因资源。     

ZmFBL41 Chang7-2: 玉米抗纹枯病的关键利器

由真菌Rhizoctonia solani引起的纹枯病严重危害玉米(Zea mays)和水稻(Oryza sativa)等作物的安全生产。R. solani的宿主范围广且抗源少, 加之相关的抗性机制研究有限, 导致纹枯病的危害长期得不到有效控制。近期, 中国科学家通过对318份玉米自交系进行全基因组关联分析, 筛选到1个与纹枯病抗性相关的、编码F-box结构域蛋白的候选基因ZmFBL41 (GRMZM2G109140)。ZmFBL41蛋白是SCF (SKP1-Cullin-F-box) E3泛素连接酶复合体的一员, 能介导复合体对肉桂醇脱氢酶ZmCAD的降解, 从而降低木质素的积累, 使玉米易感纹枯病。玉米抗病自交系Chang7-2中, 蛋白ZmFBL41 ^Chang7-2因2个关键氨基酸的变异, 不能结合并降解底物ZmCAD, 使木质素含量增加, 从而提高玉米对纹枯病的抗性。该研究率先揭示了SCF复合体可通过降解肉桂醇脱氢酶来调控植物免疫反应的新型分子机制, 为提高玉米及其它作物对纹枯病的抗性提供了重要理论依据和基因资源。     

3个玉米黄叶突变体的光合特性研究

在农业生产中光合作用是作物积累生物量的主要方式,其主要依赖于多种光合色素和完整的叶绿体结构与功能。而玉米叶色突变体对于研究叶绿体发育、提高玉米光合作用能力和产量具有重要意义。以两个玉米自交系郑58（Z58）和B73为对照,对从甲基磺酸乙酯（ethyl methanesulphonate,EMS）处理后的不同玉米诱变群体中筛选到的2株黄叶突变体yl?1（yellow leaf?1,Z58背景）、yl?2（yellow leaf?2,B73背景）以及从玉米自交系Z58中发现的1株自然黄叶突变体yl?3（yellow leaf?3）等3个表型相似的玉米黄叶突变体的形态特征、光合色素含量、叶绿素合成前体物质含量进行了比较研究。结果表明,与对照相比,3个突变体在整个生长周期内均呈现不同程度的黄叶表型、不复绿、植株矮小、发育迟缓;叶片总叶绿素、叶绿素a和叶绿素b含量均显著降低（P<0.05）,叶绿素a/叶绿素b比值显著升高（P<0.05）;不同突变体的各类叶绿素合成前体物质含量有不同程度的降低。3个突变体的黄叶表型可能是由不同基因的突变导致相关四吡咯化合物合成异常引起的。研究结果为定位引起3个突变体黄叶表型的突变基因和进一步探讨其利用潜力奠定了理论基础。     

内生真菌种类和母本基因型对内生真菌-禾草共生体叶形状和叶面积的影响

Epichloë内生真菌感染能够影响宿主植物的生长发育, 但关于内生真菌感染对宿主植物叶形状和叶面积的研究很少。该研究以羽茅(Achnatherum sibiricum)为实验材料, 采用长宽系数计算和扫描测定叶面积相结合的方法探究内生真菌种类和羽茅母本基因型对羽茅-内生真菌共生体叶形状和叶面积的影响。结果表明: 内生真菌感染与否、内生真菌种类和宿主母本基因型对反映叶形状的叶校正系数、叶片长度、宽度和长宽比均无显著影响, 经计算与验证, 确定了羽茅叶片的校正系数为0.594 9。采用该校正系数及叶长宽计算的叶面积与实测叶面积无显著差异, 且二者均未受到内生真菌感染与否、内生真菌种类或宿主植物母本基因型的显著影响。     

Quantitative Trait Locus Mapping and Candidate Gene Analysis for Plant Architecture Traits Using Whole Genome Re-Sequencing in Rice

Plant breeders have focused on improving plant architecture as an effective means to increase crop yield. Here, we identify the main-effect quantitative trait loci (QTLs) for plant shape-related traits in rice (Oryza sativa) and find candidate genes by applying whole genome re-sequencing of two parental cultivars using next-generation sequencing. To identify QTLs influencing plant shape, we analyzed six traits: plant height, tiller number, panicle diameter, panicle length, flag leaf length, and flag leaf width. We performed QTL analysis with 178 F₇ recombinant in-bred lines (RILs) from a cross of japonica rice line ‘SNUSG1’ and indica rice line ‘Milyang23’. Using 131 molecular markers, including 28 insertion/deletion markers, we identified 11 main- and 16 minor-effect QTLs for the six traits with a threshold LOD value > 2.8. Our sequence analysis identified fifty-four candidate genes for the main-effect QTLs. By further comparison of coding sequences and meta-expression profiles between japonica and indica rice varieties, we finally chose 15 strong candidate genes for the 11 main-effect QTLs. Our study shows that the whole-genome sequence data substantially enhanced the efficiency of polymorphic marker development for QTL fine-mapping and the identification of possible candidate genes. This yields useful genetic resources for breeding high-yielding rice cultivars with improved plant architecture.     

水稻条斑病抗性QTL的挖掘及相关基因的表达

条斑病是水稻(Oryza sativa)中的常见病害, 已经对我国粮食的高产稳产造成严重威胁。以典型籼稻台中本地1号与粳稻春江06的杂交F₁代花药培养双单倍体群体(DH)为材料, 用Xoc BLS256进行人工接菌, 对双亲及群体各株系的病斑长度进行测量和量化分析; 同时利用该群体业已构建的加密遗传图谱对病斑表型数据进行QTL作图分析。结果在水稻第2、4、5和8号染色体上共检测到4个效应值能区分开的QTL。对2号与5号染色体上2个较大的QTL区间内抗条斑病相关基因进行了表达分析, 结果表明这些基因在处理前后出现了不同程度的表达差异, 暗示这些基因可能是响应春江06与台中本地1号条斑病抗性差异的目标基因。研究结果为进一步克隆水稻条斑病抗性QTL奠定了重要基础。